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The Lower Permian Insects of Kansas

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The Lower Permian Insects of Kansas THE LOWER PERMIAN INSECTS OF KANSAS. PART 9. THE ORDERS NEUROPTERA, RAPHIDIODEA, CALONEURODEA AND PROTORTHOPTERA (PROBNISIDAE), WITH ADDI- TIONAL PROTODONATA AND MEGASECOPTERA." Received October 12, 1941 Presonted October 9. 1941 Dr. Tillyard's sudden deatli in 1938 left unfinished new wing is 195 min. long and 40 mln. wide (inaxi- his account of the Permian insects from the Elmo murn). This indicates that the total wing expanse limestone in the Peabody Museum at Yale Uni- was about 41 cm. or 16 inches. The holotype speci- rersity. The unstudied specimens in that collection, men, which was not quite complete, measured 165 mostly belonging to the Protorthoptera and related mm. in length.' groups, have recently been turned over to me by Tillyard's restoration of the wing of schucherti the authorities of the Peabody Museum for identifi- (1925, fig. 5) is now seen to be incorrect in two re- cation and description. The present paper deals spects. First, the wing is more slender apically than with the Caloneurodea contained in both the Yale he assumed it to be; and second, tlie subcosta ends and Harvard Collections, and with the Neuroptera, well before the apex of the wing, instead of at the Raphidiodea, and part of the Protorthoptera (family apex, as lie lias drawn it. His figure of the actual Probnisidae) in the Harvard' collection, the .Yale type specimen is incorrect in that it has lt4 + 5 specimens of these groups having already been de- (termed R3 in his paper) arising in the distal half of scribed by Tillyard. I have also included here an the wing, well beyond the level of the first fork of account of some Protodonata and Megasecoptera in MA. Actually, R4 + 5 arises (in both the holotype the Harvard collection which were not considered in and the new specimen) slightly before the middle of my earlier papers on these orders. the wing, and directly above the fork of RIB. In all except individual details tlie new wing has tlic saine Additional PROTODONATA venation as the holotype. Since the publication of my previous account of One of the most interesting aspects of the new the Protodonata from the Elmo limestone (1939), fossil is the presence of an oblique vein at the point Professor R. C. Smith, of Kansas State College, has of separation of R2 from R3. This is obviously the sent me a magnificently preserved hind wing of homologue of the vein (subnodus) described by Megatypus schucherti Tillyard (family Meganeu- Sellards in Typus permianus (1906, p. 253, figs. 4, 5). ridae). It was collected by two entomological As I have already stated (1939, p. 38) such a vein students in the College, Mr. Otto Wenger and Mr. has not previously been discernible in any Proto- Floyd Holmes; the reverse of the fossil is in the donata (Megatypus or Typus) in the Yale or Har- Kansas State College entomological collection and vard collections from Elmo. It should be noted the obverse in tlie Museum of Coinparative Zoology. that there is no indication of the oblique vein "oJ' Apart from doubtful fragments, M. schucherti lias described by Sellards in Typus. The obliquity of previously been known only from the holotype, a this vein is almost certainly an individual trait. hind wing lacking the distal fourth of the wing and some details of venation (Tillvard, 1925). The new Additional AIEGASECOPTERA bpecimen is an absolutely complete hind wing, The extinct order Afegasccoptera is not uncoin- shoa,ing with perfect clarity all veins (Plate I, inonly represented in certain Carboniferous strata figure 1). It is not only interesting as the largest of Europe, but until comparatively recently it has entire wing yet found in the Permian, but it is also not been found in Permian rocks. The insects which i~i~portantbecause it enables us to complete our Tillpard originally described from the Elmo lime- howledge of the venation of this huge insect. The stone as members of a new order, Protohymenoptera, * This investigatiorl has been aided by a grant from The width of th~holotype wing was given by Till- tile Permanent Science Fund of the American Acade~rly yard as 94 mm., but this is an error; it sl~ouldhave of Arts and Srienres. been 39 mIn. are now generally recognized as close relatives of the Genus Bltt~ouTillyarcl Megasecoptera and as constituting merely a sub- Elmoa Tillyard, 1937, Amer. Journ..Sci. 33: KX. order of the Megasecoptera. In his last discussion Fore wing: long and moderately slender; costal of the affinities of the Protohymenoptera, Tillyard space of moderate width; Sc terminating on Rl (1936) expressed agreement with this view, though before mid-wing ;Rl unbranched; pterostigma absent claiming it to be a matter of personal choice whether or at most very weakly formed; Rs with at least the group be regarded as a suborder or an order. three terminal branches; R, M and CuA separating The Protohymenoptera from the Elmo limestone at the same point; MA probably unbranched; MA contained in the Harvard collection have already joined to Rs by a weak cross-vein shortly after the been described (Carpenter, 1931a, 1931b, 1933, origin of the latter; MP forked; CuA unbranched; 1939). cross-veins few and weakly developed. Hind wing: Apart from the Protohymenoptera, the order similar to the fore wing in venation, except for dif- Megasecoptera is represented in the limestone by ferences noted under the account of the family. several species which are more closely related to the Genotype: Elmon trisecta Till. Eumegasecoptera2 than to the Protohymenoptera. One of these, Elmoa trisecta Tillyard, was recognized Elmoa trisecta Tillyard by Tillyard as a Megasecopteron and placed by him in a new family, Elmoidae. Two other species which Figure 1 he described from the formation (Martynovia insig- Elmoa ~risecta Tillyard, 1937, Amer. Journ. Sci. nis and Martynoviella protohymenoides) are, in my 33: 84. opinion, also Megasecoptera, though they were con- Fore wing: length, 12 mm. ; width, 3.2 mm. ; 3 or 4 sidered by him to be Neuroptera, belonging to the strong costal veinlets at base, above stem R+M, suborder Sialoidea (Megaloptera). Unfortunately, and 1-3 weaker ones more distal along Sc; a few not only was each of these three species based upon oblique veinlets between R1 and costal margin, a single wing, but the type of Elmoa trisecta con- especially distally; Sc terminating beyond origin of sisted of a basal fragment of a wing. Had these Rs; stem R + M arched; R4 + 5 long, arising before specimens been more nearly complete, Tillyard mid-point of Rs; R2 and R3 at most half as long as would undoubtedly have recognized the very close R4 + 5; MA arising proximally to origin of Rs; MP relationship between Elmoa, which he placed in the forked to about half its length; CUPnot forked, but Megasecoptera, and the Martynoviidae, which he connected to hind margin by a long cross-vein; 1A assigned to the Xeuroptera. The evidence for my remote from hind margin, connected to it by at conclusion on the affinities of these fossils is derived least one cross-vein; 2A very short. from a study of the specimens in the Harvard col- Hind wing: length, 12 mm.; width, 4 mm.; more lection and will be considered below, after a descrip- nearly oval than the fore wing; costal space as wide tion of the insects. as in fore wing, with similar veinlets; Sc not quite so long as in fore wing and nearer R + M than in fore Suborder EUMEG.~SECOPTERA wing. Family ELMOIDAE The body structure is preserved in one specimen (No. 4591ab); this shows only the thorax, part of Wings subequal; membrane glassy. Fore wing: the head, and a small piece of the abdomen. The Sc terminating before or at mid-wing; costal veinlets head is small (1.5 mm. long) and broad, and appar- few and weakly developed; Rs arising before mid- ently has prominent eyes. wing; M coalesced proximally with R; MA and MP Holotype: No. 15591ab, Peabody Museum, Yale present; MA not coalesced with Rs or R1; CuA University. This consists of the basal third of a coalesced proximally with stem R + M; two anals hind wing, not a fore wing, as stated by Tillyard. present ; pterostigma apparently absent. Hind In the Harvard collection there are eight speci- wing: narrow basally, but broader near the middle mens of trisecta, of which six are from the upper than the fore wing; Sc closer to R than in fore layer of the limestone: no. 4591ab, a complete insect, wing; rest of venation as in fore wing. Almost showing both pairs of wings; the fore wings overlap nothing is known of the body structure. part of the hind pair, but the two pairs are readily At present the family consists of one genus. distinguished. No. 4606ab, a very nearly complete The order Megasecoptera includes the two sub- fore wing, well preserved; no. 4595ab, 4607ab, each orders Eumegasecoptera and Pr~t~ohymenoptera(Car- consisting of the proximal third of a fore wing; no. penter, 1933). 4590ab, a complete and excellently preserved hind THE LOWER PERMIAN INSECTS OF KANSAS. PART 9 FIGURE1. Elm.oa trisecta Till. A, fore wing: drawing based mainly on specimen no. 4604, Mus. Comp. Zool. B, hind wing; drawing based chiefly on specimen no. 4590, Mus. Comp. Zool. Sc, subcosta (concave); R1, radius (convex); Rs, radial sector (concave); MA, anterior media (convex); MP, posterior media (concave); CuA, anterior cubitus (convex); CUP, posterior cubitue (concave); IA, 2A, anal veins. wing; and nos. 4592ab, 4593, 4594ab, all hind wings, which it represents was more generalized than other more or less complete.
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